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Decision making
Editorial overview
Kenji Doya and Michael N Shadlen
Current Opinion in Neurobiology 2012, 22:911–913
For a complete overview see the Issue
0959-4388/$ – see front matter, # 2012 Elsevier Ltd.
All rights reserved.
http://dx.doi.org/10.1016/j.conb.2012.10.003
Kenji Doya
Okinawa Institute of Science and
Technology, Neural Computation Unit, 1919-1
Tancha, Onna, Okinawa 904-0412, Japan
e-mail: doya@oist.jp
Kenji Doya received his PhD in 1991 at U.
Tokyo. He was a research associate at U.
Tokyo, U. C. San Diego, and Salk Institute
before joining Advanced Telecommunications
Research Institute International (ATR) in 1994.
In 2004, he was appointed as the principal
investigator of Neural Computation Unit,
Okinawa Institute of Science and Technology
(OIST) and started Okinawa Computational
Neuroscience Course (OCNC) as the chief
organizer. As OIST re-established itself as a
graduate university in 2011, he became a
professor and the vice provost for research.
He serves as the co-editor in chief of Neural
Networks from 2008. He is interested in
understanding the functions of basal ganglia
and neuromodulators based on the theory of
reinforcement learning. He is also a triathlete
and finished Ironman Hawaii in 2006 and 2011.
Michael N Shadlen
Professor of Neuroscience, Investigator,
Howard Hughes Medical Institute, Columbia
University Medical Center, 1051 Riverside
Drive, PI Kolb Annex 873, New York,
NY 10032, USA
e-mail: ms4497@columbia.edu
Michael N Shadlen received his PhD in 1985
from UC Berkeley and his MD in 1988 from
Brown University. He was professor of
physiology and biophysics at the University of
Washington until 2012. He is an Investigator of
the Howard Hughes Medical Research
Institute and professor of neuroscience at
Columbia University. Shadlen studies the
neural mechanisms that underlie decision
making, visual perception, and timing. His
studies combine neurophysiology, behavioral,
and computational techniques. Shadlen is
also a neurologist and a jazz guitarist.
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Strictly speaking, we do not make decisions, decisions make us.
– Jose Saramago, All the Names
Almost any interesting cognitive function can be framed as a decision of
some sort, because once the function admits flexibility, contingency, or a
provisional plan, it embraces elements of deliberation and commitment.
Similarly, many behaviors which we do not regard as obviously cognitive —
which proceed without conscious thought or ideation — also rely on decision
processes. Even for the most basic, existential act of waking up from sleep or
anesthesia or a minimally conscious state approximating coma, some uncon-
scious process decides to engage the world. The decision to do this in
response to the baby’s cry but not the traffic or thunder attests to the
sophistication of even this most basic operation of our mental lives.
Viewed from this perspective, the neurobiology of decision making offers a
window on to brain mechanisms that support cognition, and it allows us to
appreciate the rudiments of cognition in simpler behaviors. Thus to some,
the central importance of this area of inquiry is its exposure of principles of
cognitive neuroscience. To others, the neurobiology illuminates many
intrinsically interesting issues pertaining to decision-making itself: how
we operate as agents in a marketplace (Levy and Glimcher; Takahashi;
Rangel and Clithero; Adams et al.), society (Seo and Lee; Adams et al.), or
how we view ourselves as moral agents (Roskies).
This issue of COIN reflects a wide range of topics, models, techniques and
interests in the field of decision making. Its contributions span moral
philosophy, neuroeconomics, learning, foraging, perception and motor con-
trol in humans, nonhuman primates, and rodents. The articles draw upon
brain imaging, EEG, MEG, single neuron recordings, mathematics, and
philosophy. Although it is by no means comprehensive, there ought to be
something in this issue for just about anyone with an interest in the topic. Yet
despite this diversity, there are a variety of themes that link many of these
articles.
Several contributions focus on perception and action. This is not surprising
since sensory psychophysics has always incorporated a decision stage to
connect perception to choice and response time. The neurobiology of
perceptual decisions, as exemplified in Romo et al. and Churchland and
Ditterich, shifts the emphasis from the representation of sensory data to the
accumulation of evidence bearing on a proposition, interpretation or categ-
orization. This accumulation and a host of other factors are bundled into a
decision variable that is but a hair-trigger (or threshold crossing) away from
commitment to a choice.
Current Opinion in Neurobiology 2012, 22:911–913
912 Decision making
Most experimentally tractable decisions involve some
kind of action to indicate a choice. Indeed, much progress
in our understanding of the neural mechanisms of
decision-making derives from the study of neurons
involved in action selection, planning, or canceling.
The success of this program reminds us that cognitive
functions like decision making evolved as an elaboration
on a simpler sensory-motor design of the brain. Yet, there
is something unsettling about this: decisions do not feel
like they are about actions but about propositions. A
commitment to a proposition may be communicated
through an action, but the decision itself seems to concern
something more abstract. We see two sides of this issue
batted about by Cisek and Dehaene and Sigman. Cisek
expands on a Gibsonian theme of affordances — the idea
that even perceptual qualities are to be understood in
terms of the way we grasp, approach, sit upon, and so
forth — whereas Dehaene and Sigman argue forcefully
for the limitations of this perspective. They speculate that
neurobiology of cognitive decisions will require more
elaborate architecture than we can grasp from affordances
and plans of action. An open question is whether some-
thing qualitatively different is required — language or a
central executive that can broadcast a message to many
possible brain targets — or whether the basic mechan-
isms and architecture will look like provisional plans of
action, albeit with rule or strategy substituted for action,
what might be termed decisions about decisions about
decisions — ultimately — to act or engage or explore
elsewhere (Rushworth et al.; Botvinick).
An appealing idea is that similar computational principles
are at play in a wide variety of decisions, whether they
ultimately concern propositions, perceptions, actions,
value preferences or strategies. For propositions and per-
ceptual decisions, Bayesian theory reigns supreme, but
Bayes is not just about inference. As explained by Drugo-
witsch and Pouget, the Bayesian formulation countenances
costs and therefore has the power to link perceptual-based
and value-based decisions. Thus it is reassuring that the
formalisms in Rangel and Clithero and Adams et al. which
deal with value-based decisions should resemble those that
have proven so useful in perceptual decisions [1]. Once
costs and gains are brought into the fold, even social
decisions seem like they might involve the same types
of neural computations. Interestingly, Seo and Lee raise
the possibility that social cues can enter decision making
both as costs/gains and as contextual cues bearing on the
type of decision to engage in the first place.
The Bayesian formulation entices us to glimpse the
principles behind sublime cognitive capacities, but they
also expose parallels with the ostensibly mundane neural
processes devoted to moving our bodies (Wolpert and
Landy). It appears that the neurobiology of decision
making has more than a superficial connection to
the way the motor system chooses and implements
Current Opinion in Neurobiology 2012, 22:911–913
movement trajectories, and how the motor system
updates commands based on noisy proprioceptive and
visual feedback it receives in flight. A more obvious
decision for the motor system is to act in the first place
(Desmurget and Sirigu) and to cancel an action that has
been planned (Schall and Godlove). The contribution
from Desmurget and Sirigu describes fascinating work on
the neurobiology of volition, a topic that intersects
traditional philosophical problems of free will and moral
responsibility, as discussed in Roskies.
Whereas it seems secure to posit that many types of
decisions draw on common computational principles, it
is less clear whether the same neural processes are
involved. One possibility is that single neurons convert
all elements of a decision — evidence, value, social costs,
elapsed time — into a common currency of neural
activity. This perspective is expressed forcefully in Levy
and Glimcher based on human fMRI, and it is the subtext
of many contributions in the issue. It may be challenging
to reconcile this view with the compelling division of
functions proposed in Rushworth et al.. This article pro-
vides a wonderful example of the type of hierarchical-but-
intertwined architecture required to make decisions
about something at the moment, but to retain — for
learning or further exploration or both — unchosen
options and their associated values.
As the field uncovers computational principles, it will be
important to dig deeper into their mechanisms. How do
neural circuits give rise to computations such as integ-
ration — which underlies evidence accumulation —
valuation, setting bounds, detecting bound crossings
and so forth. An immediate challenge is to model these
processes with realistic neural elements. It is truly a
challenge because there is so little known about how
neurons or circuits of neurons can display firing rates that
reflect the integration of evidence over time scales that
are orders of magnitude greater than the membrane time
constant. Contributions from Wang and Cain and Shea-
Brown expose the challenges and potential solutions to
the neural integrator. In our view, this is one of the most
important problems in cognitive neuroscience. An overly
leaky neural integrator is the likely culprit behind the
neurology of confusion, wandering attention, poor con-
centration and many other disorders of higher brain
function. And an unstable integrator — runaway self-
excitation — may well be at the root of tics and epilepsy.
The mechanisms involved in valuation and processing
rewards may be more tractable. Clark et al. show in
rodents that there are two separate but related systems
for valuation: a mesolimbic dopamine system for reactive
decisions and an orbitofrontal system for cognitive de-
cisions. Takahashi demonstrates using PET with radio
ligands that there are links between risk-seeking and
dopamine and loss-aversion and norepinephrine. These
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same neural systems are also believed to underlie the
positive reinforcement that supports learning.
Indeed, the theory of reinforcement learning (RL) has
played an important role in many studies of decision
making. RL relies on neural computations that would
associate reward with behavioral ‘state’ and the menu of
possible actions, but as argued by Dayan the translation of
these theoretical entities into neural representations
remains unclear. Even the definition of reward, which
is arguably the most straightforward, may be nuanced
once one acknowledges that the acquisition of novel
information can be a reward in itself. While many de-
cisions are reactive and almost automatic, we often spend
time and energy deliberating about what can happen
following different options and how good or bad they
can be. Doll et al. review the dichotomy of model-based
and model-free RL and highlight enigmatic overlaps of
the neural substrates of the two systems. Rushworth et al.
review the functions of different regions of the prefrontal
cortex and suggest parallel valuation mechanisms
involved in searching and deciding.
Another theme that arises when synthesizing the variety
of topics in this issue concerns the stochastic nature of
choice. Neural mechanisms underlying perceptual de-
cisions operate deterministically, although they appear
stochastic because noise in the stimulus and brain lead to
the appearance of probabilistic behavior. This noise is real
and unavoidable, but when an animal chooses A over B on
70% of trials, it does not mean that the brain computed
P(A is better) = 0.7 and rendered a random binary out-
come that matches that probability. A proper understand-
ing of the mechanism should explain why the decision
variable corresponds to ‘A is probably better than B’ on
70% of trials. It is interesting to read contributions in this
issue and wonder whether the theories seem to imply a
matching of probabilities or the maximizing (with noise)
that is the mainstay of signal detection theory and its
extension to bounded evidence accumulation (e.g. diffu-
sion models). It is interesting to read the piece on free will
by Roskies with this same dialectic in mind.
There is much missing from this issue. Invertebrate and
rodent models of perceptual decision making are under-
represented despite rapid progress in this field [2–4]. This
issue contains little information about the mechanism for
establishing a termination criterion and sensing that this
criterion is met by neurons that represent the decision
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Editorial overview Doya and Shadlen 913
variable. This seems like a tractable problem [5,6], but
the fact is, we do not know which brain structures are
responsible. Further, we do not know how — once a
decision has terminated — neurons activate other neural
structures to initiate a response, stop the integration
process, or initiate another decision process. For example,
we do not know how contextual cues lead to a decision to
treat information about color, say, as a evidence bearing
on where to look. We might postulate mechanisms that
accumulate evidence and achieve some threshold level of
activation. But after that, what are the neural events that
bring a circuit into play and configure it so that the
relevant evidence bears on the appropriate set of possible
actions? These circuit-selection and circuit-configuration
problems are likely to involve feedback and cortico-
thalmo-cortical pathways that instantiate what might be
termed a ‘decision to engage’ [7].
We hope the articles in this issue will excite readers by
highlighting the progress of this burgeoning field and by
drawing attention to the challenges and mysteries that lie
ahead. We hope readers will delight in the insights while
yearning for deeper understanding of the neural mech-
anisms. Decision-making brings neuroscience into the
domain of ethics, philosophy and the law and thus strikes
at what it is that makes us human. And if this sounds scary,
fear not, for explaining is not the same thing as explaining
away. Rather, the neurobiology illuminates the mechan-
isms that, as Saramago put it, make us who we are.
References
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3. Uchida N, Kepecs A, Mainen ZF: Seeing at a glance, smelling in a
whiff: rapid forms of perceptual decision making. Nat Rev
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4. Harvey CD, Coen P, Tank DW: Choice-specific sequences in
parietal cortex during a virtual-navigation decision task.
Nature 2012, 484:62-68.
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Current Opinion in Neurobiology 2012, 22:911–913